RESEARCH Open Access Genetic diversity of selected genes that are potentially economically important in feral sheep of New Zealand Grant W McKenzie 1 , Johanna Abbott 2 , Huitong Zhou 1 , Qian Fang 1 , Norma Merrick 1 , Rachel H Forrest 3 , J Richard Sedcole 1 , Jonathan G Hickford 1* Abstract Background: Feral sheep are considered to be a source of genetic variation that has been lost from their domestic counterparts through selection. Methods: This study investigates variation in the genes KRTAP1-1, KRT33 , ADRB3 and DQA2 in Merino-like feral sheep populations from New Zealand and its offshore islands. These genes have previously been shown to influence wool, lamb survival and animal health. Results: All the genes were polymorphic, but no new allele was identified in the feral populations. In some of these populations, allele frequencies differed from those observed in commercial Merino sheep and other breeds found in New Zealand. Heterozygosity levels were comparable to those observed in other studies on feral sheep. Our results suggest that some of the feral populations may have been either inbred or outbred over the duration of their apparent isolation. Conclusion: The variation described here allows us to draw some conclusions about the likely genetic origin of the populations and selective pressures that may have acted upon them, but they do not appear to be a source of new genetic material, at least for these four genes. Background It is thought that livestock genetic variation has decreased through breed s ubstitution and crossing of local and global breeds [1]. Accordingly, interest in feral populations has increased because they are potential sources of genetic variation that may have been lost in commercial sheep flocks [2,3]. It has been argued that reintroducing genetic variability could enhance produc- tion in commercial breeds [4]. New Zealand (NZ) has eleven feral sheep populations either on the mainland, or on offshore islands [5]. The mainland populations originated from farmed sheep [6], while those on offshore islands either originated from farms, or were liberated as a food source for mariners [7]. These populations have been described previously [1,4,6,8-13]. In this study, the level of genetic variation of four genes was determined in order to ascertain whether th e isola- tion of these flocks had preserved greater genetic diversity compared to their commercial counterparts in NZ. These four genes are located on three different chromosomes i.e. KRTAP1-1 (chromosome 11; a keratin-associated pro- tein gene that encodes a protein KAP1-1 commonly found in wool), KRT33 (chromosome 11; encoding wool keratin K33), ADRB3 (chromosome 26; encoding the seven-transmembrane domain beta-3 adrenergic receptor ADRB3) and DQA2 (chromosome 20; encoding a class II major histocompatibility complex (MHC) protein DQA2). Previous studies have reported that variations in the keratin and keratin-associated protein g enes, including the ones above, influence many wool properties includ- ing fibre diameter [14], staple strength [15], mean staple length [16] and the brightness of wool [16]. Accordingly, * Correspondence: Jon.Hickford@lincoln.ac.nz 1 Department of Agricultural Science, Faculty of Agriculture and Life Sciences, PO Box 84, Lincoln University, Lincoln 7647, New Zealand Full list of author information is available at the end of the article McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Genetics Selection Evolution © 2010 McKenzie et al; licensee BioMed Centr al Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution Licen se (http://creativecommons.org/licenses/by/2.0 ), which permi ts unrestricted us e, distri bution, and reprodu ction in any medium, provided the original work is properly cited. given the wide phenotypic variation seen in the wool of feral sheep [6,8], one might expect to see increased var- iation in these genes. Neonatal lamb mortality, particularly in Merino sheep, represents a large loss to the NZ sheep industry. Allelic variation in ADRB3 has been associated with survival in various sheep breeds [17], thus it might be expected that previously reported or new alleles would be found at a higher frequency in feral populations routinely exposed to harsh environmental conditions. It has been reported that feral sheep may have an incr eased resistance to a number of diseases. This resis- tance could imply that variation in key immune function genessuchasthehighlypolymorphicMHCgenesis important, as it plays a role in the immune response to pathogens and parasites [18-21]. Collectively the four genes chosen here cover a variety of different animal traits that could be associated with variation in the ability to survive in remo te and poten- tially more severe environments, and where feed avail- ability was probably reduced relative to farmed sheep. Materials and methods Sheep and DNA sources Ten feral flocks and two reference flocks (non-feral) were investigated in this study (Table 1). Genomic DNA from these sheep was obtained from whole blood col- lected on FTA Classic Cards (Whatman Bio Science, Middlesex, UK) following the manufacturer’sinstruc- tions. Reference flock allele frequencies (see Tables 2 and 3) were sourced from published data [17,22-24] and from NZ commercial sheep DNA samples stored at Lincoln University. PCR amplification and genotyping PCR amplifications and ge notyping approaches were carried out using previously described methods [17,24-26] Data analysis Allele frequencies, number of alleles, observed hetero- zygosity (H O ), expected heterozygosity (H E with a Levene’s correction) and coefficient of inbre eding (F IS ) esti mates based on the method of Weir and Cockerham [27] were determined using GENEPOP version 4.0.7 [28]. This software was also used to determine devia- tions from Hardy-Weinberg equilibrium (HWE) using the Exact Test with a Markov Chain Method [29] (10 batches, 5 000 iterations per batch and a dememoriza- tion number of 10 000). Corrections for multiple sig nifi- cance tests were performed using Fisher’smethodand by applying a sequential Bonferroni type correction [29]. F IS estimates were calculated across all the populations and genes (global F IS ) and for individual populations and genes. Allelic richness, a measure of genetic diver- sity at a single locus, was determi ned using FSTAT ver- sion 2.9.3 [30] and included rarefaction to correct for sample size variation [31]. Allele frequencies for each feral population were compared to those of Merino sheep sourced from NZ Table 1 Origin of sheep populations and sample numbers (N) Flock type Flock location Origin/Breed/Type N Feral Offshore Arapawa Island I Australia/Merino/unknown 17 Arapawa Island II Australia/Merino/unknown 61 Chatham Island Australia/Merinos/Saxon 22 Pitt Island Australia/Merinos/Saxon 519 Campbell Island Australia/Merino × longwool 105 Mainland Woodstock Australia/Merino/unknown 31 Hokonui I Tasmania/Merino/Saxon 12 Hokonui II Tasmania/Merino/Saxon 73 Herbert Tasmania/Merino/unknown 24 Mohaka Unknown/Merino/unknown 14 878 Domestic reference flocks Mainland Merino 1 New Zealand/Merino/unknown 20 123 All breeds 2 Corriedale, Poll Dorset, Suffolk, Borderdale, Coopworth, Dorset Down × Coopworth, Merino × Coopworth, Merino × Polwarth, Merino, Polwarth, Dorset Down and Hampshire, NZ Romney, Awassi, Finnish Landrace and other NZ crossbred sheep 43 737 McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 2 of 8 Table 2 Within-population sample sizes (N), number of alleles identified (n) and allele frequencies for KRTAP1-1, KRT33 and ADRB3 KRTAP1-1 Population N n ABC Arapawa Island I 14 3 0.43 a 0.46 a 0.11 bc Arapawa Island II 59 2 - 0.93 c 0.07 bc Chatham Island 22 2 0.18 ab 0.82 ab - Pitt Island 477 2 - 0.85 b 0.15 a Campbell Island 97 3 0.02 c 0.82 b 0.16 a Woodstock 28 3 0.29 a 0.68 ab 0.04 b Hokonui I 11 2 - 0.82 ab 0.18 bc Hokonui II 65 2 - 0.88 b 0.12 bc Herbert 23 2 0.15 ab 0.85 b - Mohaka 6 2 - 0.83 ab 0.17 bc Merino reference flock 795 3 0.23 a 0.7 a 0.07 b All Breeds reference flock 309 3 0.06 b 0.80 b 0.14 c KRT33 Population N n ABCDE Arapawa Island I 13 5 0.04 a 0.19 ab 0.46 ac 0.19 a Arapawa Island II 60 3 - - 0.40 a 0.23 a 0.38 b Chatham Island 22 5 0.32 b 0.07 a 0.16 bc 0.23 c 0.23 a Pitt Island 471 4 0.04 a - 0.05 c 0.43 c 0.48 b Campbell Island 92 5 0.02 a 0.42 b 0.01 a 0.10 b 0.45 b Woodstock 30 5 0.07 c 0.18 c 0.30 b 0.33 c 0.12 a Hokonui I 11 4 0.23 bc 0.36 b - 0.23 bc 0.18 a Hokonui II 67 5 0.27 b 0.28 b 0.14 b 0.21 a 0.10 a Herbert 24 5 0.13 bc 0.35 b 0.31 b 0.19 b 0.02 c Mohaka 14 2 - - - 0.43 c 0.57 b Merino reference flock 739 5 0.26 b 0.36 b 0.19 b 0.04 b 0.15 a All Breeds reference flock 967 5 0.08 c 0.04 a 0.05 c 0.40 c 0.43 b ADRB3 Population N n A 1 B 2 C 3 D 4 E 1 F 2 G 5 H 5 Arapawa Island I 17 4 0.32 bc - 0.35 bc - 0.24 bc 0.09 bc Arapawa Island II 60 4 0.17 a - 0.04 a - 0.39 a 0.40 a Chatham Island 22 4 0.27 bc - 0.16 c - 0.55 a 0.02 c Pitt Island 499 6 0.20 a 0.04 a 0.23 c - 0.28 a 0.25 a 0.002 a - Campbell Island 102 4 0.66 a 0.17 a 0.1 c - - 0.01 a Woodstock 30 4 0.28 bc 0.05 bc 0.18 c - - 0.48 a Hokonui I 11 3 0.73 a 0.23 bc - - 0.05 a Hokonui II 68 4 0.53 a 0.29 a - - 0.16 bc - 0.01 a - Herbert 24 4 0.77 a 0.04 bc 0.17 c - - 0.02 b - Mohaka 6 3 0.25 bc 0.08 bc - - - 0.67 a Merino reference flock 4 484 6 0.35 b 0.02 b 0.33 b 0.06 0.20 b 0.05 b All Breeds reference flock 13 420 8 0.37 c 0.09 c 0.21 c 0.02 0.20 c 0.10 c 0.01 b 0.004 1-5 represent the effect of gene on cold survival based on the odd ratios reported in [17]: 1 good survival; 2 neutral survival; 3 below average survival; 4 poor survival, 5 data insufficient to determine the effect on survival; a-c allele frequency differences within columns that share no common alphabetic superscripts are significantly different (P < 0.05), while those pair wise comparisons that are not different are represented with the same superscripts; “-” represents alleles or data not available McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 3 of 8 commercial farms [ 17] and to the combined allele fre- quencies in breeds commonly found i n NZ [17,22-24]. This was undertaken to determine which groups were more closely r elated to each other based on “ distance” measured by the Pearson c 2 -statistic for eac h possib le pair of breeds and their respective estimated gene frequencies. Results All genes investigated in this study were polymorphic and allele frequ encies for each gene varied among the studied flocks (Table 2 and 3). No new allele was identified for any of the genes in any of the sheep typed in this study. All the KRTAP1-1 alleles previously described were pre- sent in the feral sheep except allele A absent in four breeds, including one population from Arapawa Island (Hokonui sheep) and one from Pitt Island (Mohaka sheep), and allele C absent in the populations from Herbert Forest and Chatham Island. Previous studies have reported five KRT33 alleles [25], all of which occurred in the feral popula tions. Alleles D and E were found in all the populations whereas alleles A and B were absent in the sheep from Arapawa Island and the Mo haka populations, alle le C was absent in thosefromMohakaandallelesB and C in those from the Pitt Island and Hokonui, respectively. Six different ADRB3 alleles were detected in the feral sheep. The lowest diversity was observ ed in the Mohaka population with only three alleles while it was greatest in the sheep from Pitt Islan d with six alleles. It is inter- estin g to note that alleles D and H, which occur at rela- tively low frequ encies in other commercial breeds in NZ [17], were absent in all the feral populations. The fre- quency of allele G is low in NZ commercial sheep and Table 3 Within population sample sizes (N), number of alleles identified (n) and allele frequencies for DQA2 1 DQA2 alleles Population N n 06023 0601 08011 0901 0103 1101 0102- 1601 0101- 1401 1201 Arapawa Island I 17 8 0.18 a - 0.15 a - 0.06 a 0.06 a - - 0.03 a Arapawa Island II 61 7 0.02 a 0.02 a - - - - 0.03 a 0.17 a - Chatham Island 22 6 0.23 a - - 0.05 a - 0.09 a Pitt Island 519 13 0.07 b 0.02 b - 0.11 b 0.03 b 0.02 a 0.04 a 0.14 a 0.11 b Campbell Island 105 9 - 0.01 b - - 0.07 a 0.17 b - 0.40 b - Woodstock 31 10 0.03 b - - - 0.05 a 0.31 b - - 0.24 b Hokonui I 12 6 0.05 a - 0.13 a - 0.08 a - - 0.63 b 0.04 b Hokonui II 73 9 0.07 a 0.08 a 0.21 b - 0.12 a 0.13 b - 0.23 a 0.11 b Herbert 24 7 0.06 a - 0.35 b - 0.02 b 0.08 a - 0.23 a - Mohaka 8 2 - - - - - 0.94 b - 0.06 a - Merino reference flock 20123 - - - - - - - 0.04 a All breeds reference flock 43737 - 0.11 a 0.04 a 0.03 a 0.08 a 0.10 a 0.01 a 0.05 a 0.15 a 0.13 b DQA2 alleles Population 08012- 0201 0701- 1401 0701- 1301 0401- 1501 0702- 1401 0301 0501 0402- 1701 0401- 1601 Arapawa Island I 0.32 a - - - 0.09 bc - 0.12 b Arapawa Island II 0.11 a - - - 0.07 bc - 0.58 a Chatham Island 0.27 a - - - 0.23 a - 0.14 b Pitt Island 0.002 c - - - 0.33 a 0.001 a 0.01 a 0.13 a - Campbell Island - - - - 0.16 a 0.17 a 0.005 a 0.02 bc 0.005 a Woodstock 0.05 bc 0.02 a 0.02 a 0.23 a 0.03 bc - - 0.03 bc - Hokonui I - - - - 0.08 bc - - Hokonui II - - - 0.01 c 0.05 bc - - Herbert 0.19 a - - - - - 0.06 b Mohaka - - - - - - Merino Reference flock 0.03 b - 0.001 a 0.04 b 0.03 b - - 0.02 b 0.05 b All breeds reference flock 0.003 c 0.02 a 0.002 a 0.03 c 0.02 c 0.02 b 0.06 b 0.02 c 0.16 c 1 DQA2 nomenclature [24]; a-c allele frequency differences within columns that share no common alphabetic superscripts are significantly different (P < 0.05), while the pair-wise comparisons that are not different are represented with different superscripts; “-” represents alleles or data not available McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 4 of 8 wasonlyfoundatalowfrequencyinthesheepfrom Pitt Island and Hokonui. The distribution of DQA2 alleles varied c onsiderably among popula tions with some alleles completely absent in some populations. The lowest diversity was observed in the sheep from Mohaka with only two DQA2 alleles. Conversely, thirteen DQA2 alleles were present in the sheep from Pitt Island. For all four genes, i n most cases allele frequencies in the feral populations differed significantly (p ≤ 0.025) from the frequencies in the reference flocks, most of the differences being highly significant (p < 0.001). The fol- lowing exceptions were found: (1) frequencies of KRTAP1-1 alleles of sheep from Chatham Island (p = 0.113), Woodstock (p = 0. 434), Herbert (p = 0.055) and Mohaka (p = 0.098) were not significantly different from those of the Merino reference flock, and those of sheep from Mohaka (p = 0.673), Campbell Island (p = 0.084) and Hokonui I (p = 0.454) were not different from those of all breeds and (2) frequencies of ADRB3 alleles of shee p from the Arapawa Island I did not differ from those of e ither reference flock (Merino p = 0.332; All breeds p = 0.771). Allelic richness, observed (H O ) and expected (H E ) levels of heterozygosity and coefficient of inbreeding (F IS )are shown in Table 4. On average between 2.03 and 4.86 alleles were detected per polymorphic gene across all the populations. The lowest number of alleles was observed for the ADRB3 gene (1.59) in the sheep from Arapawa Island II while the greatest number of alleles was found for KRT33 (6.12) in the Hokonui II sheep. Allelic richness was highest for KRT33 and lowest for AD RB3 in all feral populations except for the Mohaka sheep. Observed and expected heterozygosity v alues ranged from a low of 0.06 observed for KRTAP1-1 to a hig h of 1.0 for KRT33, and a low of 0.13 for KRTAP1-1 and a high of 0.86 for DQA2, respectively. Arapawa I sheep had the highest mean estimate for H O and H E over all of the genes (0.73 and 0.73, respectively), while Mohaka sheep had the lowest mean estimate for Ho and He (0.43 and 0.38, respectively). Allele sharing w as high between animals originating from Campbell Island and Pitt Island for KR TAP1-1 and among the Arapawa II flock of feral sheep for KRT33 and lower among the Arapawa I flock for DQA2. Finally, allele sharing among sheep from Woodstock was very low for DQA2. Table 4 Allelic richness (r), expected (H E ) and observed (H O ) heterozygosity, F IS 1 values for feral sheep populations of New Zealand Population Arapawa Island I Arapawa Island II Chatham Island Pitt Island locus r H O H E F IS r H O H E F IS r H O H E F IS r H O H E F IS KRTAP1-1 3.72 0.71 0.61 -0.18 3.30 0.14 0.13 -0.06 3.17 0.36 0.30 -0.20 4.24 0.23 0.26 0.11* KRT33 5.96 0.69 0.73 0.05 4.00 0.58 0.65 0.11** 5.02 0.81 0.78 -0.05 5.95 0.59 0.58 -0.02 ADRB3 2.83 0.71 0.73 0.03 1.59 0.75 0.66 -0.13 1.94 0.59 0.62 0.04 1.87 0.77 0.77 -0.01 DQA2 4.26 0.82 0.84 0.02* 2.96 0.66 0.62 -0.06 4.48 0.95 0.81 -0.18 2.83 0.82 0.83 0.00 Mean 4.19 0.73 0.73 - 2.96 0.53 0.52 - 3.65 0.68 0.63 - 3.72 0.60 0.61 - Population Campbell Island Woodstock Hokonui I Hokonui II locus r H O H E F IS r H O H E F IS r H O H E F IS r H O H E F IS KRTAP1-1 2.90 0.06 0.30 0.79** 3.41 0.36 0.47 0.24 2.54 0.36 0.31 -0.18 3.05 0.20 0.21 0.03 KRT33 4.71 0.59 0.61 0.04 5.49 0.63 0.76 0.17 4.41 1.00 0.77 -0.33 6.12 0.85 0.78 -0.09 ADRB3 2.06 0.53 0.51 -0.37 2.38 0.80 0.66 -0.21 1.97 0.55 0.44 -0.26 1.79 0.69 0.61 -0.13 DQA2 3.03 0.74 0.76 0.02 4.33 0.94 0.81 -0.16** 3.95 0.67 0.60 -0.11 4.52 0.89 0.86 -0.04 Mean 3.18 0.48 0.55 - 3.90 0.68 0.68 - 3.22 0.65 0.53 - 3.87 0.66 0.62 - Population Herbert Mohaka Allele richness averages locus r H O H E F IS r H O H E F IS KRTAP1-1 2.61 0.30 0.26 -0.16 3.00 0.33 0.30 -0.11 3.19 KRT33 5.04 0.79 0.74 -0.07 1.86 0.57 0.53 -0.09 4.86 ADRB3 1.90 0.46 0.38 -0.20 2.00 0.67 0.53 -0.29 2.03 DQA2 4.03 0.79 0.79 -0.01 2.00 0.14 0.14 - 3.64 Mean 3.40 0.59 0.54 - 2.22 0.43 0.38 - 1 Significance of F IS is indicated * P< 0.05, ** P< 0.01, figure in bold character shows a tendency towards significance (P < 0.10); negative values indicate outbreeding while positive values indicate inbreeding; “-” represents data that could not be obtained McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 5 of 8 Discussion This is the first report describing DNA varia tion in feral sheep from NZ. The genes investigated in this study were chosen because they had previously been shown to influence wool traits [16], cold survival [17] and footrot resistance [20,21]. No new allele was identified for any of the genes in the feral sheep, suggesting that they will not be a source of alternative genetic variability, at least for these genes. The allelic richness and heterozygosity results (observed and expected) are comparable with those presented in previous studies of non-NZ wild sheep populations [32-34]. Although the feral sheep sampled were chosen so that they were representative of their populations, there is no guarantee that the farmers who maintain these p opula- tions on the NZ mainland have been able to maintain genetic diversity, especially because the flocks sizes are small compared to the original populations. Allele shar- ing among four offshore island flocks (Arapawa I and II, Pitt Island and Campbell Island) was significant for one gene but not necessarily for the same gene. Sheep popu- lations from both Pitt and Campb ell Islands, have undergone extensive size reduction before being relo- cated to the mainland and it is surprising that the level of inbreeding is not higher. In contrast, among the mainland Woodstock sheep, many different alleles are detected for DQA2 suggesting this flock is o utbred, although other loci would need to be typed to confirm this. This is most likely due to the ongoing introduction of new genetic material from other Merino sheep which are typically farmed in areas adjacent to this population. Sources of genetic variation in the feral sheep popula- tions include founder effects, random drift, b alancing selection , genetic bottlenecks, or combinations of these. Each will be discussed below. Genetic drift may have affected these feral populations [35]. However, in some feral populations allele freq uen- cies were similar to those in commercially far med Mer- ino sheep. This may not be surprising since both the feral and commercial merino sheep share the same Aus- tralian origin, and the two groups have been separated at most by 50 generations. In some case s, allele frequencies in the feral popula- tions were not “Merino-like” and tended to show greater similar ity to allele frequencies in other common farmed sheep in NZ. This provides support for the anecdotal contention that these feral sheep have at times interbred with farmed non-Merino sheep. There is evidence of genetic differences between groups of sheep on remote yet neighbouring islands. Chatham and Pitt Island sheep are thought to be des- cendents of the same founding Merino sheep, yet they show quite different allele frequencies for many of the gen es studied here. Pitt and Chatham Island feral sheep have distinct wo ol colours but whether this is a result of the differences in the genes studied cannot be ascer- tained here. Founder effects may influence the genetic diversity of feral populations [36]. It is apparent from early farming records that many of these flocks were initiated with 50 or less animals and hence the likel ihood of finding rare alleles in the founding individuals might be small. Both ADRB3 variants D and H are rare in farmed NZ sheep [17] and they are absent from the feral populations stu- died here. An alternative explanation to the founder effect is that particular ADRB3 alleles have been lost in the feral populations because they provide no selective advantage. This is called balancing selection and it reflects the situation where alleles are retained in a population by forms o f selection such as heterozygote advantage, fre- quency-dependent s election [37] or selection varying in space and time that favours some alleles in certain environments [38]. ADRB3 alleles A and E are asso- ciated with cold survival, alleles C and F are linked to cold-related mortality, and allele D has a strong associa- tion with cold-related mortality and total mortali ty [17]. The complete absence of ADRB3 all ele D in the feral populations could be due to the fact t hat these flocks were exposed to cold climatic conditions during lambing and death of lambs carrying the allele. A number of studies have suggested that feral sheep show few signs of susceptibility to infection by ectopara- sites [9,12] and fly strike [39] when compared to o ther domesticated sheep breeds. The reason why these ani- malsmaybemoreresistanttoparasitesremains unknown, but may involve genetic variation or reduced/ non-exposure to the pathogens. Charbonnel and Pemberton [40] have suggested that infection with Teladorsagia circumcincta imposes a selection pressure in the Soay sheep of the island of Hirta in Scotland, and that this is refle cted in the tem- poral divergence of the MHC genes over a relatively short period between 1988 and 2000. In the context of the results reported here, while the MHC allelic richness is at times low, in the absence of any data or evidence of on-going disease challenge it would be speculative to attempt to draw any conclusions. It should be noted that for DQA2, allele sh aring was high within one island population but low within the mainland feral popula- tion, suggesting that the island population may have undergone some selection pressure. Allele sharing at KRT33 and KRTAP1-1 was typically low suggesting the flocks may be outbred. A llele rich- ness was highest for KRT33 indicating that the level of genetic diversity ha s remained quite high in these feral McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 6 of 8 sheep populations. F eral sheep populations have some unique wool characteristics including at times a hairy birth-coat type, which has been shown to offer some advantage in improving lamb survival [41-43] , the ability to shed their wool [7], tightly curled wool [12] and var- ious coat colours and markings [8]. The genes responsi- ble for these traits have yet to be identified, but may includesomeofthegenesforkeratinsandKAPsthat constitute wool fibre. Genetic bottlenecks can cause loss of genetic diversity [44]. L ike founder effects , they are largely responsible for the loss of low-frequency alleles and tend to increa se the abundance of inte rmediate- and high-f requency a lleles [45]. It is generally admitted that sheep populations from Pitt and Campbell Islands originated from a small num- ber of founding animals that multiplied subsequently. After reaching a size of ap proximately 4 000 sheep on both islands, genetic bottlenecks most likely occurred, when the majority of the sheep were slaughtered, and smal l numbers of sheep were transferred to NZ to creat e the flocks studied here. Thus these island populations may have been subject to both founde r and bottleneck effects, but the data presented her e does n ot show any strong evidence in favour of the historically documented bottlenecks and there are no obvious differences in allelic richness between the Pitt and Campbell island popula- tions compared to the other feral sheep populations. Acknowledgements This research was supported by the Brian Mason Scientific and Technical Trust. We appreciate the time and effort of the farmers who maintain these sheep and for their generosity in supplying the DNA samples. We also thank members of the Gene-Marker Laboratory for completing the genotyping of samples. Author details 1 Department of Agricultural Science, Faculty of Agriculture and Life Sciences, PO Box 84, Lincoln University, Lincoln 7647, New Zealand. 2 Environment Canterbury, PO Box 345, Christchurch 8140, New Zealand. 3 Faculty of Sport and Health Sciences, Eastern Institute of Technology, Private Bag 1201, Napier, New Zealand. Authors’ contributions GM supervised the collection of the genotype data, completed most of the analyses and drafted the manuscript. HZ and QF generated the DQA2 genotype data. NM provided the genotype data for the keratin genes used in this study. RF developed the ADRB3 genotyping methodology and generated the allele frequency data for the Merino and all breeds reference populations used in this study. She also helped revise this manuscript. JA applied for and was granted the funding that underpinned the collection of blood and data from the owners of these sheep. She designed the study and collected the blood samples from the different sheep populations identified. She was involved in typing KRTAP1-1 and assisted draft the manuscript. JRS provided completed parts of the statistical analysis and provided useful discussion on the results obtained from this study. He also assisted in the production of the final manuscript. JH helped develop the project in his capacity as research leader, provided comments on the grant proposal, and drafted the final manuscript. All authors read and approved the final manuscript. Competing interests The authors declare that they have no competing interests. Received: 1 March 2010 Accepted: 21 December 2010 Published: 21 December 2010 References 1. Tapio M, Miceikiene I, Vilkki J, Kantanen J: Comparison of microsatellite and blood protein diversity in sheep: inconsistencies in fragmented breeds. Mol Ecol 2003, 12:2045-2056. 2. Mendelsohn R: The challenge of conserving indigenous domesticated animals. 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Alexander G: Temperature regulation in the new-born lamb. IV. The effect of wind and evaporation of water from the coat on metabolic rate and body temperature. Aust J Agric Res 1962, 13:82-99. 44. Luikart G, Sherwin WB, Steele BM, Allendorf FW: Usefulness of molecular markers for detecting population bottlenecks via monitoring genetic change. Mol Ecol 1998, 7:963-974. 45. Luikart G, Allendorf FW, Cornuet JM, Sherwin WB: Distortion of allele frequency distributions provides a test for recent population bottlenecks. J Hered 1998, 89:238-247. doi:10.1186/1297-9686-42-43 Cite this article as: McKenzie et al.: Genetic diversity of selected genes that are potentially economically important in feral sheep of New Zealand. Genetics Selection Evolution 2010 42:43. Submit your next manuscript to BioMed Central and take full advantage of: • Convenient online submission • Thorough peer review • No space constraints or color figure charges • Immediate publication on acceptance • Inclusion in PubMed, CAS, Scopus and Google Scholar • Research which is freely available for redistribution Submit your manuscript at www.biomedcentral.com/submit McKenzie et al. Genetics Selection Evolution 2010, 42:43 http://www.gsejournal.org/content/42/1/43 Page 8 of 8 . RESEARCH Open Access Genetic diversity of selected genes that are potentially economically important in feral sheep of New Zealand Grant W McKenzie 1 , Johanna Abbott 2 ,. 89:238-247. doi:10.1186/1297-9686-42-43 Cite this article as: McKenzie et al.: Genetic diversity of selected genes that are potentially economically important in feral sheep of New Zealand. Genetics Selection Evolution 2010 42:43. Submit. Characteristics of some rare breeds of sheep. Rare Breeds NewZ, No.59, December 2002 (Rare Breeds Conservation Society of New Zealand;[http://www.rarebreeds.co.nz]. 9. Whitaker RH: Feral sheep in New Zealand.